Angaracris barabensis ( Pallas, 1773 )

Storozhenko, Sergey Yu., Benediktov, Alexander A. & Huang, Jianhua, 2017, Revision of the genus Angaracris Bey-Bienko, 1930 (Orthoptera: Acrididae, Oedipodinae) with notes on the tribe Bryodemini, Zootaxa 4244 (3), pp. 416-428 : 417-421

Angaracris barabensis ( Pallas, 1773)

Figs 1–16 , 24–26

Gryllus (Locusta) barabensis Pallas, 1773: 728 (syntypes—males and females, Russia: West Siberia, Barabinskaya steppe [steppes on right side of Irtysh River from Omsk to Altai]; lost).

Oedipoda barabensis: Fischer von Waldheim, 1846: 296 , Pl. 14, fig. 2.

Bryodema barabensis: Brunner von Wattenwyl, 1882: 167 ; Saussure, 1884: 181; Zubowsky, 1898: 98; Zubowsky, 1900: 17; Bolívar, 1901: 226, 233; Karny, 1908: 48; Kirby, 1910: 262; Bey-Bienko, 1925: 53; Bey-Bienko, 1929: 103.

Bryodema barabense: Jacobson, 1905: 266 ; Miram, 1907: 8; Pylnov, 1918: 138;

Angaracris barabensis: Bey-Bienko, 1930a: 119 , Pl. 18, fig. 6, Pl. 19, fig. 3; Bey-Bienko, 1930b: 498; Miram, 1933: 38; Berezhkov, 1937: 47, 69; Zimin, 1938: 73, Pl. 9, figs. 47, 49; Tarbinsky, 1948: 125; Bey-Bienko & Mistshenko, 1951: 604, figs. 1234, 1236; Berezhkov, 1951: 21; Berezhkov, 1956: 162; Bey-Bienko, 1964: 282; Popov, 1964: 1314; Popov, 1965: 122; Čejchan & Mařan, 1966: 186; Ivanova, 1967: 134; Chogsomzhav, 1972: 183; Mistshenko, 1972: 113; Mařan & Čejchan, 1974: 16; Zheng & Hang, 1974: 429; Harz, 1975: 446; Chogsomzhav, 1977: 87; Mistshenko, 1968: 495; Li, 1981: 167; Storozhenko, 1986: 317, figs. 115(12), 114(6); Sergeev, 1986: 213; Vysotskaya & Bugrov, 1987a: 1190; Chogsomzhav, 1989: 94; Kang et al., 1989: 343; Otte, 1995: 328; Benediktov, 1998: 792, figs. 1(7, 15), 2(5), 3, 4(1), 5; Li, 1998: 72, 553; Dubatolov & Sergeev, 1999: 55; Childebaev & Storozhenko, 2001: 39; Cherniachovskij, 2002: 167, 199, fig. 8.85; Sergeev & Childebaev, 2002: 297; Korsun, 2004: 93; Childebaev et al., 2013: 2, figs. 1-4; Benediktov, 2016: 469, fig. 4.

Angaracris barbensis (!): Bi, 1986: 195. Incorrect subsequent spelling.

Angeracris (!) barabensis: Steinmann, 1965: 113 ; Steinmann, 1967: 119; Steinmann, 1968: 247. Incorrect subsequent spelling.

Oedipoda hospes Fischer von Waldheim, 1846: 295 , Pl. 24, figs. 1, 2 (syntypes—male and female, Russia: Baikal Lake; lost). Synonymized with Bryodema barabensis by Saussure (1884).

Oedipoda lugubris Fischer von Waldheim, 1846: 298 , Pl. 32, figs. 4, 5 (syntypes—male and female, Russia: Daourie [= south part of Zabaikalskyi krai]; lost). Synonymized with Bryodema barabensis by Saussure (1884).

Oedipoda rhodopa Fischer von Waldheim, 1836: 348 (syntypes—males and females, Russia: Altai Mts, Katun Ridge; lost). Synonymized with Angaracris barabensis by Benediktov (1998).

Oedipoda rhodoptera (!): Fischer von Waldheim, 1836: Pl. 4, fig. 2. Incorrect subsequent spelling.

Oedipoda rhodopa: Fischer von Waldheim, 1846: 296 , Pl. 21, fig. 4.

Bryodema barabense var. rhodopus: Jacobson, 1905: 266 ; Miram, 1907: 8.

Bryodema barabense var. rhodopa: Ikonnikov, 1911: 256 .

Byodema rhodopa: Kirby, 1910: 262 .

Angaracris rhodopa: Bey-Bienko, 1930a: 121 ; Bey-Bienko, 1930b: 498; Miram, 1933: 38; Tarbinsky, 1948: 125; Bey-Bienko & Mistshenko, 1951: 604; Berezhkov, 1956: 163; Bey-Bienko, 1964: 282; Popov, 1964: 1314; Čejchan & Mařan, 1966: 186; Ivanova, 1967: 134; Mistshenko, 1968: 495; Chogsomzhav, 1972: 183; Mistshenko, 1972: 113; Mařan & Čejchan, 1974: 17; Zheng & Hang, 1974: 429; Chogsomzhav, 1977: 87; Li, 1981: 168; Yin, 1982: 84, fig. 17 (a-d); Bi, 1986: 195; Storozhenko, 1986: 317; Sergeev, 1986: 213; Vysotskaya & Bugrov, 1987a: 1190; Chogsomzhav, 1989: 94; Kang et al., 1989: 343; Zheng & Ren, 1993: 428, 429; Otte, 1995: 328; Li, 1998: 70, 553, fig. 36 (a-f).

Angaracris barabensis rhodopa: Furukawa, 1939: 66 , 121, 152.

Angaracris rhopodus (!): Berezhkov, 1937: 47. Incorrect subsequent spelling.

Angaracris rhadopus (!): Berezhkov, 1951: 21. Incorrect subsequent spelling.

Angeracris (!) rhodopa: Steinmann, 1965: 113 . Incorrect subsequent spelling.

Bryodema barabensis var. rhodoptila Karny, 1908: 49 (syntypes—sex and number of type specimens not recorded; North-West China: Gansu province, Lantschou; depositary of types unknown). Synonymized with Angaracris rhodopa by Bey-Bienko (1930).

Bryodema barabense var. roseipennis Krauss, 1901: 237 (syntypes— 9 males, 5 females; China: Kuku-nor Ridge in Nanshan Mts. [= Qilian Mts., North-West China]; in SMNS Stuttgart). Synonymized with Bryodema barabense var. rhodopus by Jacobson (1905).

Oedipoda thunbergi Stål, 1861: 345 (holotype—female, type locality unknown, wrongly labeled “ Rio Janeiro, West Indies ”; in NHRS Stockholm). Synonymized with Bryodema barabensis by Saussure (1884).

Oedipoda thunbergi: Walker, 1870: 734 .

Angaracris nigripennis Lian & Zheng, 1984: 305 , 310, figs. 23-27 (holotype—male, China: Gansu, Sunan; in IZSNU Shaanxi). Synonymized with Angaracris barabensis by Childebaev et al. (2013).

Angaracris nigripennis: Bi, 1986: 195 ; Li, 1998: 75, 553, fig. 39 (a-f); Zeng & Han, 1998: 26; Otte, 1995: 328.

Angaracris acrohylina Bi, 1986: 195 , figs. 1-5 (holotype—male; China: Tibet, Lhasa; in IEAS Shanghai). Syn. nov.

Angaracris acrohylina: Zheng & Ren, 1994: 251 , 253; Otte, 1995: 328; Li, 1998: 77, 553, fig. 40 (a-c); Zeng & Han, 1998: 26.

Angaracris morulimarginis Huang, 1981: 78 , 83, Pl. 3, fig. 6 (holotype—male, China: Tibet, Lhasa; in IZAS Beijing). Syn. nov.

Angaracris morulimarginis: Bi, 1986: 195 ; Otte, 1995: 328; Li, 1998: 74, 553, fig. 38.

Angaracris morulipennis Zheng & Ren, 1994: 251 , 253, figs. 7, 8 (holotype—male, China: Ningxia, Zhongwei; in IZSNU Shaanxi). Syn. nov.

Angaracris neimongolensis Zeng & Han, 1998: 25 , 28, fig. 1 (holotype—male, China: Inner Mongolia, Zhenglanqi; in IZSNU Shaanxi). Syn. nov.

Angaracris nigrimarginis Zheng & Ren, 1993: 427 , 429, figs. 11, 12 (holotype—female, China: Inner Mongolia, Baorou; in IZSNU Shaanxi). Syn. nov.

Angaracris ulashanicus Li, 1981: 168 , 173, figs. 1-3 (holotype—male, China: Inner Mongolia, Ulashan; in IZAS Beijing). Syn. nov.

Angaracris ulashanicus: Lian & Zheng, 1984: 306 , 310; Li, 1998: 73, 553, fig. 37 (a-d); Otte, 1995: 328.

Angaracris ulashanica (!): Bi, 1986: 195. Incorrect subsequent spelling.

Type material. Types of species described by Pallas (1773) and Fisher von Waldheim (1836, 1846) are lost. For stability of nomenclature and according to Article 75 of the Code (ICZN, 1999) we designate the neotypes for follow taxa:

Gryllus (Locusta) barabensis Pallas, 1773 . Neotype—male , Russia: “ Severnaya , Slavgorodskiy u., Omskoy gub.” [=Altaiskiy krai, Kliuchevckoy rayon, Severka, 52o08'N, 79o17'E], 7 August 1922, coll. V. Raevsky ( ZISP); here designated ( Fig. 1 ).

Oedipoda hospes Fischer von Waldheim, 1846 . Neotype—male , Russia: Irkutskaya oblast, 15 km E Ust- Ordynsky, 52o48'N, 104o46'E, 4 August 1994, coll. P.G. Nemkov ( ZISP); here designated ( Fig. 2 ).

Oedipoda lugubris Fischer von Waldheim, 1846 . Neotype—male , Russia: Zabaikalskiy krai, 35 km SW Borzya, 50o24'N, 116o33'E, 20 July 1977, coll. S.Yu. Storozhenko ( ZISP); here designated ( Fig. 3 ).

Oedipoda rhodopa Fischer von Waldheim, 1836 . Neotype—male, Russia: Altai Republic, Onguday , 50o44'N, 86o08'E, 12 July 2016, coll. M.Yu. Proshchalykin ( ZISP); here designated ( Fig. 4 ).

Other material examined. Totally 1092 specimens. ZISP (941 specimens) including 29 males & 14 females from Kazakhstan , 254 males & 221 females from Russia , 223 males & 165 females from Mongolia, and 19 males & 16 females from China ; IBPS (154 specimens) including 92 males & 53 females from Russia and 2 males & 4 females from Mongolia .

Description. Width of vertex between eyes 1.6–1.8 times wider than the width of frontal ridge between antennae; depression of vertex shallow, without median carinula. Frontal ridge relatively broad, slightly widened near the base of antennae and distinctly depressed at least near the median ocellus. Antennae reaching the hind margin of pronotum; mid segments of antennae 2.0–2.6 times as long as wide. Metazona in male 1.9–2.3 times, in female 1.8–2.1 times longer than prozona. Metasternal interspace in male 1.9–2.0 times, in female 2.3–2.4 times as broad as long. Tegmina in rest position surpassing apex of hind femur in male on 11.0– 12.2 mm and in female on 6.0– 9.5 mm; maximal width of costal field 1.4–2.0 times wider than the width of subcostal field and almost equal to the width of medial field. Male supra-anal plate with two widely rounded ridges near the base and two almost parallel ridges in apical part ( Fig. 7 ). Male subgenital plate conical ( Fig. 8 ). Female basivalvular plates rugulose ( Fig. 10 ).

Coloration of body, wings and legs is extremely variable. Body brownish, greenish or blackish, usually with black and/or white spots especially on pronotum. Tegmina brown, greenish or blackish, with numerous dark spots or stripes, rarely without any marks. The color of hind wings is also extremely variable from hyaline membrane to provide by dark spots, stripes and bands as well as to the completely black or dark brown membrane ( Fig. 16 ). The base of anal field of the hind wings yellowish, greenish or rose; anal veins blackish, yellowish, brownish or red. Hind femora pale brown or greenish, rarely almost completely black; outer side of femur with 2–3 black bands and one light pregenicular band; lower side of femur yellowish, brown or red; inner side of femur pale brown or reddish with 2–3 large black spots which sometimes fused and forming longitudinal stripe. Hind tibia yellow, light brown or reddish.

Measurements (mm). Length of body: male 22.0–31.0, female 27.0–35.0; pronotum: male 5.5–7.2, female 5.8–8.5; tegmen: male 24.0–33.0, female 23.5–32.0; hind femur: male 11.0–14.0, female 13.0–18.0; ovipositor 2.4–2.5.

Distribution. Kazakhstan (north and central part: Kostanay region, Akmola region, Pavlodar region, Kent Mountains, Karsakpay Plateau), Russia (south part of Ural and Siberia: Orenburgskaya oblast, Omskaya oblast, Novosibirskaya oblast, Kemerovskaya oblast, Altaisky krai, Altai Republic, south part of Krasnoyarskiy krai, Republic of Khakassia, Tyva Republic [= Tuva], Irkutskaya oblast, Republic of Buryatia, Zabaikalskiy krai), Mongolia (Uvs aimag, Zavkhan aimag, Gov aimag, Khovsogol aimag, Bayankhongor aimag, Bulgan aimag, Ovorkhangay aimag, Dongov aimag, Omnogov aimag, Dornogo aimag, Tov aimag, Khentiy aimag, Dorod aimag), China (Qinhai province, Gansu province, Shaanxi province, Shanxi province, Heilongjiang province, Nei Mongol Zizhiqu, Xizang Zizhiqu).

Notes. While the presence of two color forms of hind wings occurs in some species of Oedipodinae , for example in Mioscirtus wagneri (Eversmann, 1849) ( Huang et al., 2013) , but during long time the hyaline and reddish forms of Angaracris was considered as distinct species or subspecies. The presence of transitional forms, the similarity in the acoustic signals of males, absence of differences in the structure of the male genitalia, and the same localization of C-heterochromatic blocks in chromosomes show that A. rhodopa is synonym of A. barabensis ( Benediktov, 1998) .

In 1981–1998 seven new species of Angaracris was described from North and Central China, each based on a single holotype or a few (3–4) type specimens. These species were again distinguished mainly by the color and pattern of hind wings. The most different forms with hyaline, reddish and almost completely black membrane of hind wings have been synonymized recently (Childebaev et al., 2013). The investigation of variability of the hind wings color and pattern in the tribe Bryodemini is made by the second author and it was found that A. barabenis is one of the most variable species ( Benediktov, 2016). Here we considered other six Chinese species as synonyms of A. barabenis .

Angaracris barabensis is abundant in steppes and semi-deserts. It overwinters in egg stage. The egg pod is described by Zimin (1938) and Cherniachovskij (2002). Nymphs occurs from mid May to June, adults—from end June to September.

Angaracris barabensis damage the pasture vegetation ( Bey-Bienko & Mishchenko, 1951; Mistshenko, 1972), but most preferable are lichen Parmelia vagans Nyl. and vascular plants Dodartia orientalis L., Artemisia terraalbae Krash. , A. austriaca Jacq. , A. schrenkiana Ledeb. , Limonium gmelini (Willd.) , Salsola sp., and Trifolium hybridum L. (Childebaev et al., 2013). Moreover, the adults and larvae of A. barabensis in the steppe prefer the areas covered by lichen Parmelia vagans . It is noteworthy that the variegated color of adults and larvae of A. barabensis allows them to be invisible among the lichens, which is very important for the survival of this species because these large insects are potential victims for the birds (Childebaev et al., 2013).

Figures

FIGURE 16. Angaracris barabensis, variability of the hind wing pattern (after Benediktov, 2016).

FIGURES 1 – 4. Angaracris barabensis, male body, dorsal view. 1, neotype of Gryllus (Locusta) barabensis Pallas, 1773; 2, neotype of Oedipoda hospes Fischer von Waldheim, 1846; 3, neotype of Oedipoda lugubris Fischer von Waldheim, 1846; 4, neotype of Oedipoda rhodopa Fischer von Waldheim, 1836. Scale bars 10 mm.

FIGURES 5 – 10. Angaracris barabensis. 5, male sternal plate, ventral view; 6, female sternal plate, ventral view; 7, male apex of abdomen, dorsal view; 8, male apex of abdomen, lateral view; 9, female apex of abdomen, lateral view; 10, female apex of abdomen, ventral view. Scale bars 2 mm.

FIGURES 11 – 15. Angaracris barabensis, male. 11, epiphallus, dorsal view; 12, the same, frontal view; 13, the same, lateral view; 14, genitalia (epihallus removed), lateral view; 15, the same, dorsal view. Ae — ancorae; Ap — apical valves of penis; Apd — apodemes; Be — bridge; Bp — basal valves of penis; Cv — valves of cingulum; Ecm — ectophalic membrane; Enm — endophalic membrane; Le — lophi of epiphallus; Os — oval sclerite; Psp — posterior projections of epiphallus; Zyg — zygoma. Scale bars 1 mm.

FIGURES 17 – 32. Bryodemini. 17 – 19, Bryodemella (Bryodemella) holdereri (Krauss, 1901), male: 17, apex of hind femur, dorsal view; 18, mesosternum, ventral view; 19, apex of hind femur, lateral view; 20, Bryodema gebleri (Fischer von Waldheim, 1836), the base of male hind tibia, dorsal view; 21 – 23, Uvaroviola multispinosa Bey-Bienko, 1930, male: 21, apex of hind femur, dorsal view; 22, base of hind tibia, dorsal view; 23, apex of hind tarsus, dorsal view; 24 – 26, Angaracris barabensis (Pallas, 1773), male: 24, base of hind tibia, dorsal view; 25, apex of hind tarsus, dorsal view; 26, apex of hind femur, lateral view; 27 – 28, Bryodemacris uvarovi Bey-Bienko, 1930, male: 27, apex of hind femur, dorsal view; 28, pronotum, lateral view; 29, Bryodemella (Marikovskiella) semenovi (Ikonnikov, 1911), male mesosternum, ventral view; 30, Bryodema lustiosum (Stoll, 1813), male pronotum, lateral view; 31, Bryodema nigripennis Mistshenko et Gorochov, 1989, female sternal plate, ventral view; 32, Andrea gorochovi Mistshenko, 1989, female sternal plate, ventral view. (Figs. 17 – 25, 27 – 30 after Benediktov, 1998; Figs. 31 – 32 after Gorochov et al., 1986).